The composition of lignin in the outer cell-wall layers of spruce and poplar has been studied and the data obtained have been compared with those of the mature reference wood in which the secondary cell wall predominates. Materials with exclusively or predominantly outer cell-wall layers were examined. Accurate data relating to the lignin monomer composition and the number of β-O-4´ bonds were obtained from pure middle lamella/primary cell wall lignin. Firstly, a 10 000 year old white spruce material, with most of the secondary cell wall missing, was studied. The aged lignin was composed of guaiacyl units only, and was slightly more condensed but otherwise similar to the reference lignin. Secondly, the developing xylem of a Norway spruce clone was analyzed during a growth season. In spring and early summer, growth is very rapid and the intention was to sample tissues in which the secondary cell-wall layers had not yet lignified, but where the outer layers at least had started to lignify. Microscopy, Klason lignin and carbohydrate analyses showed that the lignin in the developing xylem of samples from mid-June was located exclusively in the middle lamella. The lignin was more condensed, was composed of guaiacyl units only and contained more end-groups than the reference Norway spruce wood. Thirdly, the cambial tissues of a Balsam poplar clone were surveyed during a growth season. Both the phloem side and the xylem side of the cambial region were examined. The Klason lignin content and carbohydrate monomer distribution showed that in June and August the tissues on the phloem side contained material with mainly middle lamella/primary walls. In June, the xylem side in the cambial region contained mainly middle lamella/primary walls, and in August the secondary cell wall carbohydrates were being deposited. Both tissues contained lignin that was more condensed and had more end-groups than the reference lignin. In mid-June, the developing xylem had a ratio of syringyl to guaiacyl units of 0.6, whereas the ratio for the reference wood was 1.3. In the final study, lignin from the primary cell walls from a hybrid aspen cell suspension culture was investigated. The lignin contained only guaiacyl units which were more condensed than those observed in the reference poplar wood. / <p>QC 20100920</p>

Lignocellulosic plants are the most abundant source of terrestrial biomass and are one of the potential sources of renewable energy that can replace the use of fossil fuels. For a country such as Sweden, where the forest industry accounts for 10% of the total export, there would be large economical benefits associated with increased biomass yield. The availability of research on wood development conducted in conifer tree species, which represent the majority of the forestry in Sweden, is limited and the majority of research has been conducted in model angiosperm species such as Arabidopsis thaliana. However, the large evolutionary distance between angiosperms and gymnosperms limits the possibility to identify orthologous genes and regulatory pathways by comparing sequence similarity alone. At such large evolutionary distances, the identification of gene similarity is, in most cases, not sufficient and additional information is required for functional annotation. In this thesis, two high-spatial resolution datasets profiling wood development were processed; one from the angiosperm tree Populus tremula and the other from the conifer species Picea abies. These datasets were each published together with a web resource including tools for the exploration of gene expression, co-expression and functional enrichment of gene sets. One developed resource allows interactive, comparative co-expression analysis between species to identify conserved and diverged co-expression modules. These tools make it possible to identifying conserved regulatory modules that can focus downstream research and provide biologists with a resource to identify regulatory genes for targeted trait improvement. / Lignocellulosa är den vanligast förekommande källan till markburen biomassa och är en av de förnybara energikällor som potentiellt kan ersätta användningen av fossila bränslen. För ett land som Sverige, där skogsindustrin som står för 10 \% av den totala exporten, skulle därför en ökad produktion av biomassa kunna ge stora ekonomiska fördelar. Forskningen på barrträd, som utgör majoriteten av svensk skog är begränsad och den huvudsakliga forskningen som har bedrivits på växter, har skett i modell organismer tillhörande gruppen gömfröiga växter som till exempel i Arabidopsis thaliana. Det evolutionära avståndet mellan gömfröiga (blommor och träd) och nakenfröiga (gran och tall) begränsar dock möjligheten att identifiera regulatoriska system mellan dessa grupper. Vid sådana stora evolutionära avstånd krävs det mer än att bara identifiera en gen i en modellorganism utan ytterligare information krävs som till exempel genuttrycksdata. I denna avhandling har två högupplösta experiment som profilerar vedens utveckling undersökts; ett från gömfröiga träd Populus tremula och det andra från nakenföriga träd (barrträd) Picea abies. Datat som behandlats har publicerats tillsammans med webbsidor med flera olika verktyg för att bland annat visa genuttryck, se korrelationer av genuttryck och test för anrikning av funktionella gener i en grupp. En resurs som utvecklats tillåter interaktiva jämförelser av korrelationer mellan arter för att kunna identifiera moduler (grupper av gener) som bevaras eller skilts åt mellan arter över tid. Identifieringen av sådana bevarade moduler kan hjälpa att fokusera framtida forskning samt ge biologer en möjlighet att identifiera regulatoriska gener för en riktad förbättring av egenskaper hos träd.

This thesis deals with the influence of microclimatic conditions (air temperature near tree/shrub top, temperature of the stem and root zone) on the wood phenology of trees (Picea abies) and shrubs (Pinus mugo) growing on the same site in the alpine tree line (ATL) ecotone. Phenological and microclimatic research was carried out in the Giant Mountains at 1,370 m a. s. l. in the 2017 vegetation period, with the aim to compare the course of wood formation depending on the microclimate, to determine the temperature thresholds of wood growth and differences in the morphology of trees and shrubs. Based on the microclimate analysis of trees and shrubs, individual variables were determined (average, average minimum, average maximum, daily amplitude of air, stem and root zone temperatures) characterizing the growing season. The collection of micro-drills of trees and shrubs took place from April to October in an interval of 6 - 11 days, which were subsequently processed and examined in the laboratory. The following phenophases of wood development were distinguished: cambial, enlarging, wall-thickening, mature cells and cells of the previous ring. Subsequently, differences in the number of cells of individual phenophases were identified between trees and shrubs. Furthermore, the temperature thresholds for...

This thesis describes aspects of durability of Swedish softwood in above ground applications. The thesis consists of a summary of nine papers, which all aim to explain the existing variation in above ground microbial durability and moisture sensitivity of Norway spruce (Picea abies) and Scots pine (Pinus sylvestris). The methods used were two field tests, one accelerated durability test, and three water permeability tests. The main conclusions made in the study were that Norway spruce sapwood is more susceptible to discolouring fungi than heartwood. With respect to Scots pine, the only important factor for its natural durability above ground was whether the samples consisted of heartwood or of sapwood. There was also no systematic variation between pine heartwood from different stand origins in above ground conditions. Origin had no influence on durability in practice for neither spruce nor pine. Furthermore, in contradiction with traditional thinking, annual ring width and density had no influence on durability in any of the tests. The material came from two different samplings. The ‘old’ material was sampled in the beginning of the 1980s and consisted of Scots pine and Norway spruce from three different origins: the north, the central area, and the south of Sweden. The material was subjected to different handling conditions and surface and end-grain treatments. The ‘old’ material was used in a large above ground field test, carried out between 1985 and 1994. Unfortunately, the spruce sapwood and heartwood was not separated in this field test. In the field trial with the ‘old’ material, the most important factor for the durability of Norway spruce was the samples’ surface- and end-grain treatment. It was also observed that untreated spruce showed better durability than samples that were painted but without proper end-grain treatment. Samples originating from one of the stands were more sensitive towards air-drying than samples from the other two, which was reflected in a higher average moisture content and mass loss for the untreated samples. With respect to Scots pine, the most important factor was whether the sample consisted of heartwood or of sapwood. The heartwood samples were durable irrespective of their previous treatment. The Scots pine sapwood samples, on the other hand, had a very fluctuating moisture content if they were not surface- and end-painted. This was also reflected in higher mass losses in these samples. Even when properly surface- and end grain treated, the sapwood samples did not perform as well as the heartwood samples. The ‘new’ material consisted of Scots pine taken from six different stands, and Norway spruce from five different stands, all from areas in southern Sweden. The sampling was performed in order to achieve a large variation in wood properties. Thus, logs from areas with different climate and growth conditions were collected. This material was used for laboratory tests, and also for the second field test, which was evaluated after a test period of two years. In the second field test untreated Scots pine and Norway spruce samples from the ‘new’ material were investigated. Effects of origin and different tree diameters were examined. Furthermore, Scots pine was separated to heartwood and sapwood, and Norway spruce to mature and juvenile wood. The Norway spruce samples were sawn with vertical or horizontal annual rings. For the Norway spruce in the second field test, vertical annual rings were shown to be very beneficial for avoiding crack formation, while samples with horizontal annual rings displayed a large number of cracks. Despite this, the larger number of cracks did not result in a higher moisture content or more fungal discoloration. Juvenile wood had a slightly higher average discolouring fungal growth grading than the other spruce samples. For Scots pine, the only factor of importance was whether the samples consisted of heartwood or of sapwood. Annual ring width, density and origin had no significant effect on either fungal growth or moisture uptake. This observation held for both Norway spruce samples and Scots pine samples. An accelerated test was run in order to investigate Norway spruce sapwood and heartwood with respect to their differences in durability towards discolouring fungi and moisture uptake. The Mycologg method was used to accelerate fungal growth during a number of forced moisture cycles. The results showed that sapwood was much more sensitive to discolouring fungi than heartwood. Sapwood also recorded a higher average moisture content than heartwood. The differences that were observed between the types of samples were not due to annual ring width, density or origin. It was observed that Norway spruce sapwood samples and heartwood samples showed differences in liquid water permeability. This observation was made especially evident in a droplet absorption test. Water droplets were absorbed much faster on the sapwood samples in comparison with the heartwood samples. This higher affinity to water suggested that a sapwood surface would reach a higher moisture content than a heartwood surface. This observation provides a major explanation of why the sapwood samples showed poorer durability towards discolouring fungi in the Mycologg trials.

This thesis describes methods to obtain value–addedcompounds from TMP-turpentine obtained from the spruce, Piceaabies. The methodology focuses on biotransformations using twoapproaches: an oxygenation approach (i.e. oxygenation ofterpene hydrocarbons by cell cultures) and an esterificationapproach (i.e. lipase-catalysed transesterification of vinylacetate with terpene alcohols, and a further fractionation ofthe TMP-turpentine). The main constituents of the turpentine, a-pinene, b-pineneand limonene, were subjected to a P. abies suspension culture.Allylic oxidation formed the major products for α-pineneand β-pinene, which were further oxidised to theirrespective aldehyde or ketone. One of the minor products froma-pinene, cis-verbenol, was not only transformed into verbenonebut also isomerised to trans-verbenol. Limonene gavelimonene-(1,2)-epoxide as the major product. Fractionation of monoterpenes is accomplished throughphysical separation methods, chromatography and distillation,and lipase-catalysed transesterification of vinyl acetate withterpene alcohols. The esters of myrtenol and trans-pinocarveolwere separated from the more slowly reacting alcohols such asborneol and carveol by use of a combination of the Mucor mieheilipase and Candida antarctica lipase A as catalysts.Furthermore, the non-reacting tertiary terpene alcohols wereseparated from the reacting alcohols in a single step byCandida antarctica lipase A. Lipase-catalysed (Candida antarctica lipase B andPseudomonas cepacia lipase) transesterification of vinylacetate with sterically hindered secondary alcoholsunexpectedly yielded hemiacetals or hemiacetal esters. Thereaction conditions required to obtain these side products havebeen studied. <b>Keywords:</b>Picea abies, Pinaceae, Essential oilscomposition; Terpene alcohol; Hemiacetal; Hemiacetal ester,TMP-turpentine; Monoterpene; α-Pinene; β-Pinene;Limonene; Verbenol; Pinocarveol; Borneol; Myrtenol; Suspensioncell culture; Biotransformation; Lipase-catalysed; Oxidation;Allylic oxidation; Transesterification; Autoxidation;Separation.

No description available.

Wood is a major construction material that is used in many contexts, and for different purposes. Serious problems may arise, however, when moisture related deformations as twist occur in wood used in different types of building structures, joinery and furniture. Twist can be explained to a great degree by the helical deviation of the grain angle in relation to the longitudinal direction of the log or the sawn board. Wood fibres form a spiral within the tree, and this is a natural occurrence that is named spiral grain. The wood fibres close to the pith in Norway spruce form a left-handed spiral. In most trees the grain angle turns over to be right-handed with time. Sawn timber that exhibits large grain angles lead to problems of shape stability and stiffness in finished constructions. In this thesis the spiral grain in Norway spruce (Picea abies (L.) Karst.) was stated as well as the effect on sawn timber. The material was based on sample trees from Sweden and Finland. Samples were taken in twenty-two stands at different heights in tree. From six stands studs were sawn and dried for measuring twist and other deformations. The spiral grain was measured with the method scribe test on 390 log discs taken at the top-end of the logs. Account was given concerning changes in grain angle from pith to bark, regarding both increasing annual ring numbers and distance from pith. The development of grain angle over tree age was utilized to study whether annual growth, size of tree, height in tree as well as silvicultural treatments affected spiral grain. Moreover, the relation between grain angle and distance to pith (in mm) was used to forecast twist in sawn timber. The left-handed grain angle was at its greatest between the fourth and eighth annual rings. Thereafter for most trees the grain angle turned from left-handed to right-handed in a linear fashion, in a manner that was unique for each individual tree. The pattern of spiral grain differed significantly between different stands, regarding change of inclination with increasing age or distance from pith. The culmination of the grain angle close to the pith occurred at somewhat higher age higher up in the trunk. The grain angle decreased faster in top logs than it did in the butt logs. The largest trees within a stand had a grain angle that turned to right in a slower way than smaller ones. The thinning strength and type of thinning regime also affected the character of spiral grain in the remaining trees in a stand. There was an indication that strong thinnings, where fast growing trees are retained, may lead to more individuals in a stand that exhibit high grain angles under bark. With knowledge of the size and direction of the grain angle under bark, and the diameter of the log, calculations can be made that show how twisted the sawn timber will be after drying. This can be used for deciding whether an individual log can profitably be sawn and processed further or not. The grain angle under bark can be used to remove trees showing the greatest degree of spiral grain already in the first thinning. Silvicultural methods aiming at even and dense Norway spruce stands, which normally is practised in Scandinavia, will probably result in timber with relatively low risk concerning large grain angle and subsequent risk for twist in sawn wood.

From studies of the circadian clock in the plant model species Arabidopsis (Arabidopsis thaliana), a number of important properties and components have emerged. These include the genes CIRCADIAN CLOCK ASSOCIATED 1 (CCA1), GIGANTEA (GI), ZEITLUPE (ZTL) and TIMING OF CAB EXPRESSION 1 (TOC1 also known as PSEUDO-RESPONSE REGULATOR 1 (PRR1)) that via gene expression feedback loops participate in the circadian clock. Here, we present results from ectopic expression of four Norway spruce (Picea abies) putative homologs (PaCCA1, PaGI, PaZTL and PaPRR1) in Arabidopsis, their flowering time, circadian period length, red light response phenotypes and their effect on endogenous clock genes were assessed. For PaCCA1-ox and PaZTL-ox the results were consistent with Arabidopsis lines overexpressing the corresponding Arabidopsis genes. For PaGI consistent results were obtained when expressed in the gi2 mutant, while PaGI and PaPRR1 expressed in wild type did not display the expected phenotypes. These results suggest that protein function of PaCCA1, PaGI and PaZTL are at least partlyconserved compared to Arabidopsis homologs, however further studies are needed to reveal the protein function of PaPRR1. Our data suggest that components of thethree-loop network typical of the circadian clock in angiosperms were present beforethe split of gymnosperms and angiosperms.

Against the background of past, recent and future climate change, the present thesis addresses elevational shifts of alpine treelines in the Swedish Scandes. By definition, treeline refers to the elevation (m a.s.l.) at a specific site of the upper trees of a specific tree species, at least 2 m tall. Based on historical records, the first part of the thesis reports and analyzes the magnitude of treeline displacements for the main trees species (Betula pubescens ssp. czerepanovii, Picea abies and Pinus sylvestris) since the early 20th century. The study covered a large and heterogeneous region and more than 100 sites. Concurrent with temperature rise by c. 1.4 °C over the past century, maximum treeline advances of all species amount to about 200 m. That is virtually what should be predicted from the recorded temperature change over the same period of time. Thus, it appears that under ideal conditions, treelines respond in close equilibrium with air temperature evolution. However, over most parts of the landscape, conditions are not that ideal and treeline upshifts have therefore been much smaller. The main reason for that discrepancy was found to be topoclimatic constraints, i.e. the combined action of geomorphology, wind, snow distribution, soil depth, etc., which over large parts of the alpine landscape preclude treelines to reach their potential thermal limit. Recorded treeline advance by maximum 200 m or so over the past century emerges as a truly anomalous event in late Holocene vegetation history. The second part of the thesis is focused more on long-term changes of treelines and one specific and prevalent mechanism of treeline change. The first part of the thesis revealed that for Picea and Betula, treeline shift was accomplished largely by phenotypic transformation of old-established stunted and prostrate individuals (krummholz) growing high above the treeline. In obvious response to climate warming over the past century, such individuals have transformed into erect tree form, whereby the treeline (as defined here) has risen. As a means for deeper understanding of this mode of positional treeline change, extant clonal spruces, growing around the treeline, were radiocarbon dated from megafossil remains preserved in the soil underneath their canopies. It turned out that Picea abies in particular may attain almost eternal life due to its capability for vegetative reproduction and phenotypic plasticity. Some living clones were in fact inferred to have existed already 9500 years ago, and have thus persisted at the same spot throughout almost the entire Holocene. This contrasts with other tree species, which have left no living relicts from the early Holocene, when they actually grew equally high as the spruce. Thereafter they retracted by more than 300 m in elevation supporting that also on that temporal scale, treelines are highly responsive to climate change. The early appearance of Picea in the Scandes, suggests that Picea “hibernated” the last glacial phase much closer to Scandinavia than earlier thought. It has also immigrated to northern Sweden much earlier than the old-established wisdom. The experiences gained in this thesis should constitute essential components of any model striving to the project landscape ecological consequences of possible future climate shifts.

Die Anatomie des Holzes stellt eine entscheidende Einflussgröße bei einer Vielzahl holztechnologischer Prozesse dar. Der Beitrag soll mit Hilfe rasterelektronenmikroskopischer Bildtafeln den anatomischen Bau des Holzes der Gattung Picea anschaulich vermitteln.